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Eragrostis (Poaceae): Monophyly and Infrageneric Classification Amanda L

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Eragrostis (Poaceae): Monophyly and Infrageneric Classification Amanda L Aliso: A Journal of Systematic and Evolutionary Botany Volume 23 | Issue 1 Article 44 2007 Eragrostis (Poaceae): Monophyly and Infrageneric Classification Amanda L. Ingram Cornell University, Ithaca, New York Jeff .J Doyle Cornell University, Ithaca, New York Follow this and additional works at: http://scholarship.claremont.edu/aliso Part of the Ecology and Evolutionary Biology Commons, and the Plant Sciences Commons Recommended Citation Ingram, Amanda L. and Doyle, Jeff .J (2007) "Eragrostis (Poaceae): Monophyly and Infrageneric Classification," Aliso: A Journal of Systematic and Evolutionary Botany: Vol. 23: Iss. 1, Article 44. Available at: http://scholarship.claremont.edu/aliso/vol23/iss1/44 Aliso 23, pp. 595–604 ᭧ 2007, Rancho Santa Ana Botanic Garden ERAGROSTIS (POACEAE): MONOPHYLY AND INFRAGENERIC CLASSIFICATION AMANDA L. INGRAM1,2 AND JEFF J. DOYLE L. H. Bailey Hortorium, Cornell University, 228 Plant Science, Ithaca, New York 14853, USA 1Corresponding author ([email protected]) ABSTRACT Eragrostis is a large genus in subfamily Chloridoideae of Poaceae. Recent phylogenetic analyses have suggested that the genus may not be monophyletic, that some of its segregate genera may be better placed within Eragrostis, and that current infrageneric classifications may not represent mono- phyletic groups. We have used molecular sequence data from the plastid locus rps16 and the nuclear gene waxy from a broad sample of Eragrostis species and representatives of six of the seven segregate genera to address these issues. We found that Eragrostis is monophyletic with the inclusion of several of the segregates, including Acamptoclados, Diandrochloa, and Neeragrostis. The placement of Cla- doraphis and Stiburus is uncertain. Thellungia does not belong in Eragrostis and is actually more closely related to Sporobolus. These data also suggest that existing infrageneric classifications are inadequate and do not correspond to monophyletic groups within Eragrostis. Key words: Acamptoclados, Cladoraphis, Diandrochloa, Eragrostis, Neeragrostis, Pogonarthria, rps16, Stiburus, Thellungia, waxy. INTRODUCTION genus, see Van den Borre and Watson 1994). Since the orig- inal description there has been little agreement as to which Eragrostis Wolf is a genus of approximately 350 species species actually belong in the genus and how they are related in Poaceae (Watson and Dallwitz 1992). It is the largest ge- to each other. Species in Acamptoclados Nash, Cladoraphis nus in subfamily Chloridoideae, a group comprising about Franch., Diandrochloa De Winter, Eragrostiella Bor, Neer- 1500 species (Van den Borre and Watson 1997). Members agrostis Stiburus Thellungia of Eragrostis generally are characterized by paniculate inflo- Bush, Stapf, and Stapf have rescences, multi-floreted spikelets, glabrous three-nerved been included in Eragrostis at various times. Clayton and Renvoize (1986) also suggested a close relationship between lemmas, ciliate ligules, and C4 photosynthesis. The genus is morphologically and anatomically diverse, however, and ex- Eragrostis and Pogonarthria Stapf. hibits a wide range of variation in many characters. For in- The various modes of spikelet disarticulation that are seen stance, the panicles range from very loose and open to high- in the genus have been the most common source of char- ly contracted spicate structures (Watson and Dallwitz 1992). acters for delimiting infrageneric groups. Spikelets may dis- NAD-ME, PCK-like, and intermediate forms of leaf blade articulate from the top, from the bottom, or as a unit. Ad- anatomy are found in the genus (Van den Borre and Watson ditionally, whether the paleas are retained on the rachilla or 1994). Several major types of the bicellular microhairs com- fall with the lemmas can be an important character, as can mon to Chloridoideae are found in Eragrostis, including the the persistence of the rachilla. These characters are quite chloridoid type, the panicoid type, the Pappophorum type, variable in Eragrostis and can be seen in a number of dif- and intermediates (Prendergast et al. 1986; Amarasinghe and ferent combinations. Unfortunately, these characters are not Watson 1990). Eragrostis species range throughout the generally as useful as one might hope. There may be tem- world’s tropical and subtropical regions, and they are most poral variation that is not always obvious at particular stages commonly found in weedy disturbed areas and in dry hab- in the life cycle (e.g., the paleas may be retained slightly itats. Most of the species are of little economic importance, longer than the lemmas but are still deciduous). From a prac- but one species (E. tef; tef) is cultivated as a major cereal tical standpoint, this means that herbarium material may be crop in Ethiopia. This species is also an important forage impossible to score or misleading if not collected at precisely grass, as are several other species, including E. cilianensis the correct stage. These intermediacies also cloud the dis- and E. curvula. Due in part to its large size and wide geo- tinction of the character states. The infrageneric classifica- graphic distribution, there has been no comprehensive tax- tions proposed by Koch (1848) and numerous later botanists onomic treatment of Eragrostis, and there has been some relied heavily on these characters (see Van den Borre and debate in the recent literature as to whether the genus is Watson 1994). More recent classifications have included monophyletic (Van den Borre and Watson 1997; Hilu and some other morphological characters, such as spikelet shape Alice 2001) and how infrageneric groups should be delim- (Lazarides 1997), pubescence on the palea keels, panicle ited (reviewed in Van den Borre and Watson 1994). branching, lemma keel and margin shape and curvature, and Eragrostis was first described by Wolf (1776) from ma- floret fertility (Cope 1998). terial of E. minor (for a detailed taxonomic history of the A recent phylogenetic analysis of anatomical and mor- phological data from 56 species of Eragrostis and two seg- 2Present address: Wabash College, PO Box 352, Crawfordsville, regate genera by Van den Borre and Watson (1994) led the Indiana 47933, USA. authors to conclude that divisions based on these spikelet 596 Ingram and Doyle ALISO disarticulation characters did not represent natural groups. doraphis is a genus of two species distributed in southern They instead suggested that the genus could be divided into Africa. The plants are halophytic or glycophytic and have two subgenera based on a number of correlated anatomical free pericarps and a unique ‘‘armed’’ growth form. Panicle and morphological characters. However, no uncontradicted branches in Eragrostis species terminate with spikelets, but synapomorphies were identified for these groups, and there the terminal spikelets in Cladoraphis are lacking. This in are numerous exceptions to most of the characters proposed combination with hardened panicle branches and tough, in- as diagnostic of the subgenera. The results from the phylo- rolled leaves produces thorny appendages. Diandrochloa, a genetic analyses in their study may also have been affected genus of seven widely distributed species, was separated by the frequent occurrence of allopolyploidy in the genus. from Eragrostis by De Winter (1960) on the basis of mem- Approximately 69% of the species in the genus are poly- branous ligules (other Eragrostis species have a ligule that ploids (Hunziker and Stebbins 1986), and many of the taxa is a fringe of hairs; Watson and Dallwitz 1992). Phillips included in the Van den Borre and Watson (1994) study are (1982) rejected this treatment, however, and included these known allopolyploids. Taxa of hybrid origin can exhibit species in Eragrostis. Eragrostiella comprises five species unique combinations of morphological and anatomical char- in southern Asia and Australia and was separated from Er- acter states and when included in phylogenetic analyses can agrostis by Bor (1940) based on the spicate inflorescences produce misleading hypotheses of relationships (McDade and tufted habit found in these species. Neeragrostis was 1990, 1992). described by Bush (1903) based on a single dioecious spe- The monophyly of Eragrostis has been brought into ques- cies (ϭ N. reptans). Two additional species of Eragrostis, tion by recent phylogenetic analyses of subfamily Chlori- one dioecious and another with hermaphrodite flowers (E. doideae. Van den Borre and Watson (1997) analyzed a data hypnoides), have been positioned in Neeragrostis by some set of anatomical and morphological characters from all 166 authors. All three species are prostrate annuals distributed in chloridoid genera (as recognized by Watson and Dallwitz the Americas. Stiburus, a small genus of two species from 1992). The authors used the two subgenera of Eragrostis southern Africa, was first described by Stapf (1900). These identified in their 1994 analysis of the genus as terminals species were separated from Eragrostis on the basis of their for their phylogenetic analyses and found that they were not villous spikelets, but Phillips (1982) pointed out that a num- sister taxa and were in fact found in two widely divergent ber of Old World Eragrostis species have pubescent spike- clades. Hilu and Alice (2001) conducted a phylogenetic anal- lets and submerged Stiburus into Eragrostis. The monotypic ysis of sequence data from the plastid locus matK from 74 genus Thellungia (ϭ E. advena) is native to Australia and species in 56 chloridoid genera. Included in the sample were was first
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